317 lines
37 KiB
XML
317 lines
37 KiB
XML
<document id="7670113F003DD9A766D44CB11A0360B8" ID-CLB-Dataset="34778" ID-DOI="http://dx.doi.org/10.3897/AfrInvertebr.57.8416" ID-GBIF-Dataset="eff5b73c-570a-405a-bbee-7a273e426612" ID-Pensoft-Pub="2305-2562-1-53" ID-ZooBank="9AE415AB300C4129A21BA33590DC5B42" ModsDocAuthor="" ModsDocDate="2016" ModsDocID="2305-2562-1-53" ModsDocOrigin="African Invertebrates 57 (1)" ModsDocTitle="The first scarabaeid beetle (Coleoptera, Scarabaeidae, Melolonthinae) described from the Mesozoic (Late-Cretaceous) of Africa" checkinTime="1465882667365" checkinUser="pensoft" docAuthor="Woolley, Christopher" docDate="2016" docId="82D7B5DA139E76360A127C5CDCCD2F6A" docLanguage="en" docName="AfrInvert 57(1): 53-66" docOrigin="African Invertebrates 57 (1)" docSource="http://dx.doi.org/10.3897/AfrInvertebr.57.8416" docTitle="Ceafornotensis Woolley, 2016, gen. n." docType="treatment" docUuid="B6C43D03-2660-4A2C-BB53-BE9FF6501D79" docUuidSource="ZooBank" docVersion="8" lastPageNumber="55" masterDocId="306B706BFFEE4069960DFF867E79FFC4" masterDocTitle="The first scarabaeid beetle (Coleoptera, Scarabaeidae, Melolonthinae) described from the Mesozoic (Late-Cretaceous) of Africa" masterLastPageNumber="66" masterPageNumber="53" pageNumber="54" updateTime="1701378385594" updateUser="ExternalLinkService">
|
||
<mods:mods id="7690D99877ED7A385F08296E00B81714" xmlns:mods="http://www.loc.gov/mods/v3">
|
||
<mods:titleInfo id="2302968F7312609F5C1CCAD46DA38E0A">
|
||
<mods:title id="BF2B2D1FCBC6516AA212E8793E20BBED">The first scarabaeid beetle (Coleoptera, Scarabaeidae, Melolonthinae) described from the Mesozoic (Late-Cretaceous) of Africa</mods:title>
|
||
</mods:titleInfo>
|
||
<mods:name id="4F199661CFF04C561F55624C0959DD29" type="personal">
|
||
<mods:role id="EB90C5D2CA4EF25921551EC72AF16612">
|
||
<mods:roleTerm id="34E7297464492A7AC90E5D2C60791877">Author</mods:roleTerm>
|
||
</mods:role>
|
||
<mods:namePart id="86E3BF065B08FAC092C0A1BB5A44EDF9">Woolley, Christopher</mods:namePart>
|
||
</mods:name>
|
||
<mods:typeOfResource id="E41218E110A01BE19C6E75085282ED63">text</mods:typeOfResource>
|
||
<mods:relatedItem id="2B46B4421844478476F66AE277C3CE7A" type="host">
|
||
<mods:titleInfo id="0E2CB215FAD0A22835D54865BA4384FE">
|
||
<mods:title id="6C365A761721EB2021EBAC3A559F68D8">African Invertebrates</mods:title>
|
||
</mods:titleInfo>
|
||
<mods:part id="2935D289B0034E22EBC74F1842DDB6AD">
|
||
<mods:date id="53701363EFB5CB5D7A59CDF898DF9154">2016</mods:date>
|
||
<mods:detail id="711A801BE7462C1C7B18F2ABC3A881E6" type="volume">
|
||
<mods:number id="1954703C564E6361A8E79AB5651AB15F">57</mods:number>
|
||
</mods:detail>
|
||
<mods:detail id="9143A33255C20AFB575AAF9DA400BC0D" type="issue">
|
||
<mods:number id="01CFD877D6E56713301E1F17A8980C52">1</mods:number>
|
||
</mods:detail>
|
||
<mods:extent id="FF1AB782459DFB23389864AF07F21F57" unit="page">
|
||
<mods:start id="86C8C9D183A944D1261F01EFE9CFF3C9">53</mods:start>
|
||
<mods:end id="4E9C888EE07AA239BF473513C6D427C9">66</mods:end>
|
||
</mods:extent>
|
||
</mods:part>
|
||
</mods:relatedItem>
|
||
<mods:location id="39136690A7B5C27B892259C7B74AF887">
|
||
<mods:url id="09EE4EDF5E9E40BB4787A2946CB56B4E">http://dx.doi.org/10.3897/AfrInvertebr.57.8416</mods:url>
|
||
</mods:location>
|
||
<mods:classification id="00E546E7CEE832D2E7AC44E21AF364F2">journal article</mods:classification>
|
||
<mods:identifier id="D4EFC2ED8BBD866644A3C4BAC7BE59DA" type="DOI">http://dx.doi.org/10.3897/AfrInvertebr.57.8416</mods:identifier>
|
||
<mods:identifier id="A6CFC29FB3E3002AD599C8B9B1BC8E24" type="Pensoft-Pub">2305-2562-1-53</mods:identifier>
|
||
<mods:identifier id="E5DDFAB452669273BDD5C1D5C26E92A0" type="ZooBank">9AE415AB300C4129A21BA33590DC5B42</mods:identifier>
|
||
</mods:mods>
|
||
<treatment id="82D7B5DA139E76360A127C5CDCCD2F6A" ID-GBIF-Taxon="127878992" LSID="urn:lsid:zoobank.org:act:B6C43D03-2660-4A2C-BB53-BE9FF6501D79" httpUri="http://treatment.plazi.org/id/82D7B5DA139E76360A127C5CDCCD2F6A" lastPageId="4" lastPageNumber="55" pageId="1" pageNumber="54">
|
||
<subSubSection id="FAF9E78B12BA2760F6DF98B6C957F302" pageId="1" pageNumber="54" type="multiple">
|
||
<paragraph id="3CC080575FEB4E01641A6337669B3A6C" pageId="1" pageNumber="54">Taxon classification Animalia Coleoptera Scarabaeidae</paragraph>
|
||
</subSubSection>
|
||
<subSubSection id="4A1F8355C09E2C5A851954F27A7706A8" pageId="1" pageNumber="54" type="nomenclature">
|
||
<paragraph id="931A0B7881158524C259D52583B02F9A" pageId="1" pageNumber="54">
|
||
Genus
|
||
<taxonomicName id="82CDCF93CFB4189E69E01AA1E32A7A16" ID-CoL="62KDC" LSID="http://zoobank.org/B6C43D03-2660-4A2C-BB53-BE9FF6501D79" class="Insecta" family="Scarabaeidae" genus="Ceafornotensis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Ceafornotensis" order="Coleoptera" pageId="1" pageNumber="54" phylum="Arthropoda" rank="genus">Ceafornotensis</taxonomicName>
|
||
<taxonomicNameLabel id="DEA0032015F229E6E0D708D2066A04A9" pageId="1" pageNumber="54">gen. n.</taxonomicNameLabel>
|
||
</paragraph>
|
||
</subSubSection>
|
||
<subSubSection id="FDBF551C6BCE43FE55B2319BD5A07B0A" pageId="1" pageNumber="54" type="etymology">
|
||
<paragraph id="118310D921720AE11BAC2CBB8878EC7F" pageId="1" pageNumber="54">Etymology.</paragraph>
|
||
<paragraph id="9397A633C02DB11C1831B43586F1FB5E" pageId="1" pageNumber="54">From Old English ceafor (chafer) and Greek notius (south).</paragraph>
|
||
</subSubSection>
|
||
<subSubSection id="988983085AEABB45D236A05DE69E25C7" pageId="1" pageNumber="54" type="type species">
|
||
<paragraph id="06AE33ECE400B870B9E4CF5DD3E69896" pageId="1" pageNumber="54">Type species.</paragraph>
|
||
<paragraph id="7DCA53CCED2844D7E78C463713E0DF07" pageId="1" pageNumber="54">
|
||
<taxonomicName id="CC9DEF8B7E7D4BBACEFCD03AEBBA040F" class="Insecta" family="Scarabaeidae" genus="Ceafornotensis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Ceafornotensis archratiras" order="Coleoptera" pageId="1" pageNumber="54" phylum="Arthropoda" rank="species" species="archratiras">Ceafornotensis archratiras</taxonomicName>
|
||
sp. n.
|
||
</paragraph>
|
||
</subSubSection>
|
||
<subSubSection id="167C2647618A161104D74DDD63E5F3FF" pageId="1" pageNumber="54" type="diagnosis">
|
||
<paragraph id="1E95389FD9C19CA07925AD9E712E4DCF" pageId="1" pageNumber="54">Diagnosis.</paragraph>
|
||
<paragraph id="E7D1BD2A4D0B36E09F2B8DFAC1F5B519" pageId="1" pageNumber="54">
|
||
The placement of the fossil beetle within the superfamily
|
||
<taxonomicName id="CB082B705F4B7AD2BD6A13D8B63724C1" lsidName="Scarabaeoidea" pageId="1" pageNumber="54" rank="superfamily" superfamily="Scarabaeoidea">Scarabaeoidea</taxonomicName>
|
||
is based on the following synapomorphies as referred to in
|
||
<bibRefCitation id="DADD4FB27BF9983765AE72827663DBDD" author="Krell, F-T" journalOrPublisher="Coleopterists Society Monograph" pageId="8" pageNumber="61" pagination="120 - 143" title="Fossil record and evolution of Scarabaeoidea (Coleoptera: Polyphaga)." volume="5" year="2006">Krell (2006)</bibRefCitation>
|
||
; antennae with a lamellate club; pronotum and forelegs adapted for burrowing; protibae dilated apically.
|
||
</paragraph>
|
||
</subSubSection>
|
||
<subSubSection id="CC85C7C070E208C93CE4648E823AC3D0" lastPageId="4" lastPageNumber="57" pageId="1" pageNumber="54" type="description">
|
||
<paragraph id="6802C3FA123579E3439004225E3B1C2B" pageId="1" pageNumber="54">Character comparison.</paragraph>
|
||
<paragraph id="24D65BD6EA04906C90ED901CCAD069AF" pageId="1" pageNumber="54">
|
||
The specimen differs from
|
||
<taxonomicName id="DE4EF44350F334976CF86A6890A101CD" family="Glaresidae" lsidName="" pageId="1" pageNumber="54" rank="family">Glaresidae</taxonomicName>
|
||
,
|
||
<taxonomicName id="5CE20BDA90B8D34EAEB2B35A690609C6" family="Lucanidae" lsidName="" pageId="1" pageNumber="54" rank="family">Lucanidae</taxonomicName>
|
||
,
|
||
<taxonomicName id="CC7AA448358ACAC754063B78E2A8ACCA" family="Passalidae" lsidName="" pageId="1" pageNumber="54" rank="family">Passalidae</taxonomicName>
|
||
,
|
||
<taxonomicName id="2ACE313E15EEFA0CC359ECF024C92701" family="Trogidae" lsidName="" pageId="1" pageNumber="54" rank="family">Trogidae</taxonomicName>
|
||
by six visible sternites; from
|
||
<taxonomicName id="8AEAA978E6E4B7FE6FBE5BBDB3ECECD7" family="Diphyllostomatidae" lsidName="" pageId="1" pageNumber="54" rank="family">Diphyllostomatidae</taxonomicName>
|
||
,
|
||
<taxonomicName id="4A86ECB02CDE6DE09F943ED8C80B4CC2" family="Ochodaeidae" lsidName="" pageId="1" pageNumber="54" rank="family">Ochodaeidae</taxonomicName>
|
||
by partially division of the eye by a genal lobe, apical spurs of the tibae not serrate; from
|
||
<taxonomicName id="FC63668131B93EE18823C66247D6BE0F" family="Pleocomidae" lsidName="" pageId="1" pageNumber="54" rank="family">Pleocomidae</taxonomicName>
|
||
by developed mouthparts; from
|
||
<taxonomicName id="87A36E557FBA3CA33E314ECAE8E85CA8" family="Glaphyridae" lsidName="" pageId="1" pageNumber="54" rank="family">Glaphyridae</taxonomicName>
|
||
by broad pronotum (not subquadrate), elytra long in form, covering most of the abdomen; from
|
||
<taxonomicName id="FC23E44E6189A8C6FB9EF0FAF50411E3" family="Bolboceratidae" lsidName="" pageId="1" pageNumber="54" rank="family">Bolboceratidae</taxonomicName>
|
||
,
|
||
<taxonomicName id="9BD68B2EE725C37739FF6F803450EA47" family="Geotrupidae" lsidName="" pageId="1" pageNumber="54" rank="family">Geotrupidae</taxonomicName>
|
||
,
|
||
<taxonomicName id="991F9F45EFE5A6F1223700B87C8C185B" family="Glaphyridae" lsidName="" pageId="1" pageNumber="54" rank="family">Glaphyridae</taxonomicName>
|
||
,
|
||
<taxonomicName id="0A2A2A57B96DB010D11075F6654C246C" family="Hybosoridae" lsidName="" pageId="1" pageNumber="54" rank="family">Hybosoridae</taxonomicName>
|
||
by protibia with a single tooth to outer margin, tibia outer margin not serrate, parameres long relative to basal piece and symmetrical to subsymmetrical, median lobe absent or reduced. Additionally, left mandible broad and dentate, not sickle shaped, the form unlikely to accommodate a well-developed mesal brush, antennae lamellae without cupule, would omit the specimen from the
|
||
<taxonomicName id="9148948873AAF3D6F0D73FB2A135C959" family="Hybosoridae" lsidName="" pageId="1" pageNumber="54" rank="family">Hybosoridae</taxonomicName>
|
||
to which the habitas of the specimen otherwise closely resembles.
|
||
</paragraph>
|
||
<paragraph id="3A45F24559CD41F327F4CEAF7C35FE5B" pageId="1" pageNumber="54">
|
||
Of the subfamilies of the
|
||
<taxonomicName id="9C3E7C57AB5E30165509702B29BC96C6" family="Scarabaeidae" lsidName="" pageId="1" pageNumber="54" rank="family">Scarabaeidae</taxonomicName>
|
||
, the specimen differs from
|
||
<taxonomicName id="4802B6D62956F35AA95399DAFCB94646" lsidName="" pageId="1" pageNumber="54" rank="subfamily" subfamily="Scarabaeinae">Scarabaeinae</taxonomicName>
|
||
, by two spurs on the metatibia, middle coxae contiguous, scutellum well developed; from
|
||
<taxonomicName id="710A1DF62DF78F72A7464650CEFC69C8" lsidName="" pageId="1" pageNumber="54" rank="subfamily" subfamily="Aphodiinae">Aphodiinae</taxonomicName>
|
||
,
|
||
<taxonomicName id="1FAF1667CFBCF02BC0D93A358C2CF98E" lsidName="" pageId="1" pageNumber="54" rank="subfamily" subfamily="Allidiostomatinae">Allidiostomatinae</taxonomicName>
|
||
,
|
||
<taxonomicName id="61BC3AA576A7404EA9B6969AA31F0B21" lsidName="" pageId="1" pageNumber="54" rank="subfamily" subfamily="Orphninae">Orphninae</taxonomicName>
|
||
by spurs close-set, not separated by basal tarsomere; from
|
||
<taxonomicName id="FB4C685980BE23AE895069A4FF92829D" lsidName="" pageId="1" pageNumber="54" rank="subfamily" subfamily="Cetoniinae">Cetoniinae</taxonomicName>
|
||
by short clypeus, not restricted in width before eyes, mandibles broad and dentate, the form unlikely to accommodate a well-developed mesal brush; from
|
||
<taxonomicName id="8C9CCAC7AB7FBE2F4201DC3668E730FA" lsidName="" pageId="1" pageNumber="54" rank="subfamily" subfamily="Rutelinae">Rutelinae</taxonomicName>
|
||
by metatarsal claws symmetrical, small, apical tarsomere not enlarged and lengthened; from
|
||
<taxonomicName id="ACEF6239C26307D24D207C98C5D9DF57" lsidName="" pageId="1" pageNumber="54" rank="subfamily" subfamily="Dynastinae">Dynastinae</taxonomicName>
|
||
by pronotum and clypeus simple, lacking major protuberances, foveae or carinae.
|
||
</paragraph>
|
||
<paragraph id="7A5D8867124EEDBDAC202625C53BC743" lastPageId="2" lastPageNumber="55" pageId="1" pageNumber="54">
|
||
The presence of minor pronotal or cephalic armature cannot be discounted and pronounced dimorphic features are not present in all tribes (e.g.
|
||
<taxonomicName id="BB219D7B4C7EEF620D60ABF8980DD20B" lsidName="" pageId="1" pageNumber="54" rank="tribe" tribe="Cyclocephalini">Cyclocephalini</taxonomicName>
|
||
) or may exhibit intra-specific variation. From the existing fossil record and divergence
|
||
<pageBreakToken id="1C45183DAA9FF5C2B8AA95221B186528" pageId="2" pageNumber="55" start="start">time</pageBreakToken>
|
||
estimates (
|
||
<bibRefCitation id="C957AC7E2FDD6DFC4D9E3C7B066A28F4" author="Ahrens, D" journalOrPublisher="Proceedings of the Royal Society of London B: Biological Sciences" pageId="7" pageNumber="60" title="The evolution of scarab beetles tracks the sequential rise of angiosperms and mammals." url="10.1098/rspb.2014.1470" volume="281" year="2014">Ahrens et al. 2014</bibRefCitation>
|
||
;
|
||
<bibRefCitation id="5B0762B11B3FBFF08E16E17D37271C1A" author="Krell, F-T" journalOrPublisher="Coleopterists Society Monograph" pageId="8" pageNumber="61" pagination="120 - 143" title="Fossil record and evolution of Scarabaeoidea (Coleoptera: Polyphaga)." volume="5" year="2006">Krell 2006</bibRefCitation>
|
||
), the
|
||
<taxonomicName id="9E089ABAF3C60BB5DB679C4802D678A4" lsidName="" pageId="2" pageNumber="55" rank="subfamily" subfamily="Dynastinae">Dynastinae</taxonomicName>
|
||
are thought to have diverged from the
|
||
<taxonomicName id="18D1EC2E27362991C8D8059E2B66994D" lsidName="" pageId="2" pageNumber="55" rank="subfamily" subfamily="Rutelinae">Rutelinae</taxonomicName>
|
||
during the Palaeogene. On the balance of available evidence the placement of the specimen within the
|
||
<taxonomicName id="66D812F25166750BBF65DF23C942EA05" lsidName="" pageId="2" pageNumber="55" rank="subfamily" subfamily="Dynastinae">Dynastinae</taxonomicName>
|
||
is thought unlikely.
|
||
</paragraph>
|
||
<paragraph id="3438B60758DE280050392B892B06FD12" pageId="2" pageNumber="55">
|
||
From Mesozoic taxa the specimen differs from
|
||
<taxonomicName id="08870824BFBB88211C9AE1E54F2DB65F" lsidName="" pageId="2" pageNumber="55" rank="subfamily" subfamily="Cretoscarabaeinae">Cretoscarabaeinae</taxonomicName>
|
||
,
|
||
<taxonomicName id="8B8D9EECB3FDA7F22E050F3E01EF2D85" family="Lithoscarabaeidae" lsidName="" pageId="2" pageNumber="55" rank="family">Lithoscarabaeidae</taxonomicName>
|
||
,
|
||
<taxonomicName id="6FCEE054A892D60F66700D4E85F4E8E0" lsidName="" pageId="2" pageNumber="55" rank="tribe" tribe="Holcorobeini">Holcorobeini</taxonomicName>
|
||
(
|
||
<bibRefCitation id="8B0A2D46165C18C90B1BDD41830E8C06" author="Nikolajev, GV" journalOrPublisher="Paleontological Journal" pageId="8" pageNumber="61" pagination="513 - 520" title="Pleurostict Lamellicorn Beetles (Coleoptera, Scarabaeidae) from the Lower Cretaceous of Transbaikalia." volume="32" year="1998">Nikolajev 1998</bibRefCitation>
|
||
) and
|
||
<taxonomicName id="9CCF71039C244355D0FEA702D88E7122" class="Insecta" family="Scarabaeidae" genus="Cretorabaeus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Cretorabaeus" order="Coleoptera" pageId="2" pageNumber="55" phylum="Arthropoda" rank="genus">Cretorabaeus</taxonomicName>
|
||
Nikolajev, 1995 (
|
||
<bibRefCitation id="37E555E73CA5D3AD4B83E2A4F4693F1E" author="Nikolajev, GV" journalOrPublisher="Paleontological Journal" pageId="8" pageNumber="61" pagination="185 - 191" title="A new subfamily of Lamellicorn beetles (Coleoptera, Scarabaeidae) from the Lower Cretaceous of Transbaykalia and its phylogenetic relationships." volume="29" year="1995">Nikolajev 1995</bibRefCitation>
|
||
) by meso-, metatibia bearing a single transverse carina; from the Mesozoic genera of the tribe
|
||
<taxonomicName id="A0A524A5C920D08F267ECAC00E017392" lsidName="" pageId="2" pageNumber="55" rank="tribe" tribe="Sercini">Sercini</taxonomicName>
|
||
;
|
||
<taxonomicName id="E2C6BAD603D94E112CD18A603D5EB10C" class="Insecta" family="Scarabaeidae" genus="Lithanomala" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Lithanomala" order="Coleoptera" pageId="2" pageNumber="55" phylum="Arthropoda" rank="genus">Lithanomala</taxonomicName>
|
||
Nikolajev, 1992 and
|
||
<taxonomicName id="8D198EEC2C37A53062CEC1605C4E6B0D" class="Insecta" family="Scarabaeidae" genus="Cretoserica" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Cretoserica" order="Coleoptera" pageId="2" pageNumber="55" phylum="Arthropoda" rank="genus">Cretoserica</taxonomicName>
|
||
Nikolajev, 1998 (
|
||
<bibRefCitation id="F6BC3C3E59F3F44BFF6C34A3FEDDDECD" author="Nikolajev, GV" journalOrPublisher="Paleontological Journal" pageId="8" pageNumber="61" pagination="513 - 520" title="Pleurostict Lamellicorn Beetles (Coleoptera, Scarabaeidae) from the Lower Cretaceous of Transbaikalia." volume="32" year="1998">Nikolajev 1998</bibRefCitation>
|
||
) by apical spurs close-set, not separated by basal tarsomere. Nikolajev erected the family
|
||
<taxonomicName id="162844621DA3D966FFD2A0DCD4A54715" lsidName="" pageId="2" pageNumber="55" rank="subfamily" subfamily="Cretomelolonthinae">Cretomelolonthinae</taxonomicName>
|
||
(
|
||
<bibRefCitation id="5CD335EB9FCF74766E9BE5D6127E52E6" author="Nikolajev, GV" journalOrPublisher="Paleontological Journal" pageId="8" pageNumber="61" pagination="513 - 520" title="Pleurostict Lamellicorn Beetles (Coleoptera, Scarabaeidae) from the Lower Cretaceous of Transbaikalia." volume="32" year="1998">Nikolajev 1998</bibRefCitation>
|
||
;
|
||
<bibRefCitation id="4C6521F1F3E53DA9D4BB52D6AFE247B6" author="Nikolajev, GV" journalOrPublisher="Kazak Universiteti, Almaty" pageId="8" pageNumber="61" title="Mezozoyskiy Etap Evolyutsii Plastinchatousykh (Insecta: Coleoptera: Scarabaeoidea)." year="2007">Nikolajev 2007</bibRefCitation>
|
||
) based on four specimens of
|
||
<taxonomicName id="3A2B0CCECBE846E02D318FBECD2D5719" class="Insecta" family="Scarabaeidae" genus="Cretomelolontha" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Cretomelolontha transbaikalica" order="Coleoptera" pageId="2" pageNumber="55" phylum="Arthropoda" rank="species" species="transbaikalica">Cretomelolontha transbaikalica</taxonomicName>
|
||
Nikolajev, 1998, from the Baissa locality, Lower Cretaceous. Although some characters are common to the Orapa specimen; spur at mid and hind tibial apices close-set, mid and hind tibae with one transverse carina, the Baissa specimen possesses features of the wing which cannot be corroborated; wings are not preserved in the Orapa specimen although the form and humeral development would suggest that the beetle was capable of flight. Other characters could be considered plesiomorphic. The Orapa specimen differs by broader tarsomeres relative to the width of the tibial apices, and the longer length of the parameres relative to the basal piece (membrane).
|
||
</paragraph>
|
||
<paragraph id="C093E28E240588732B374AC034601CAD" pageId="2" pageNumber="55">
|
||
The paucity of apomorphies (
|
||
<bibRefCitation id="9343472AA0639EA12B15BB7BDF7C0826" author="Browne, J" journalOrPublisher="Systematic Entomology" pageId="7" pageNumber="60" pagination="307 - 326" title="Evolution of the scarab hindwing articulation and wing base: a contribution toward the phylogeny of the Scarabaeidae (Scarabaeoidea: Coleoptera)." url="10.1046/j.1365-3113.1998.00059.x" volume="23" year="1998">Browne and Scholtz 1998</bibRefCitation>
|
||
), and that the phylogeny of the
|
||
<taxonomicName id="544C38E54D165F0255A18DA18F0AB179" lsidName="" pageId="2" pageNumber="55" rank="subfamily" subfamily="Melolonthinae">Melolonthinae</taxonomicName>
|
||
is considered paraphyletic (
|
||
<bibRefCitation id="17D13C2F5EB6AEE2AEFF85C84EBF612D" author="Ahrens, D" journalOrPublisher="Systematic Entomology" pageId="7" pageNumber="60" pagination="113 - 144" title="The phylogeny of Sericini and their position within the Scarabaeidae based on morphological characters (Coleoptera: Scarabaeidae)." url="10.1111/j.1365-3113.2005.00307.x" volume="31" year="2006">Ahrens 2006</bibRefCitation>
|
||
) and unresolved (
|
||
<bibRefCitation id="66AC5D00C8DBBA914E05C83830BA5751" author="Jameson, ML" editor="Arnett, Jr RH" journalOrPublisher="CRC Press LLC, Boca Raton" pageId="8" pageNumber="61" pagination="1 - 5" title="Series Scarabaeiformia Crowson 1960 (= Lamellicornia), Superfamily Scarabaeoidea Latreille 1802, Introduction." volumeTitle="American Beetles, Volume I: Archostemata, Myxophaga, Adephaga, Polyphaga: Staphyliniformia" year="2000">Jameson and Ratcliffe 2000</bibRefCitation>
|
||
) makes the placement of fossils into this subfamily problematic. However, lack of support for the
|
||
<normalizedToken id="DE1514EC69F8BD9E38853A0FD21F92E1" originalValue="specimen’s">specimen's</normalizedToken>
|
||
inclusion outside of the 'higher
|
||
<normalizedToken id="82F8B60F634942298560DF9DCA055BDD" originalValue="scarabs’">scarabs'</normalizedToken>
|
||
together with relative length of the parameres to the basal piece (Fig. 4c), elongate and narrow antennal lamellae and evidence from the fossil record, give support for the
|
||
<normalizedToken id="E25A9CA6ED8623F37586DEF84CCAEFA3" originalValue="specimen’s">specimen's</normalizedToken>
|
||
placement within the
|
||
<taxonomicName id="642297872405A52A2EBE0A31DD60DDA4" lsidName="" pageId="2" pageNumber="55" rank="subfamily" subfamily="Melolonthinae">Melolonthinae</taxonomicName>
|
||
.
|
||
</paragraph>
|
||
<paragraph id="79A141A829AAA85DDD0EE639371C748C" pageId="2" pageNumber="55">
|
||
Systematic phylogeny: Phylogenetic analysis was performed using the character set from
|
||
<bibRefCitation id="0D69B1E872383C13B63AD24A339341D7" author="Bai, M" journalOrPublisher="Acta Geologica Sinica" pageId="7" pageNumber="60" pagination="984 - 993" title="Prophaenognatha, a new Aclopinae genus from the Yixian Formation, China and its phylogenetic position based on morphological characters (Coleoptera: Scarabaeidae)." url="10.1111/j.1755-6724.2011.00532.x" volume="5" year="2011">Bai et al. (2011)</bibRefCitation>
|
||
. Included in the analysis are 20 species comprising 12 families of the
|
||
<taxonomicName id="FD6F6AD676D0FAE9E17ADAC908ABB92C" lsidName="Scarabaeoidea" pageId="2" pageNumber="55" rank="superfamily" superfamily="Scarabaeoidea">Scarabaeoidea</taxonomicName>
|
||
, a fossil specimen from Yixian Formation,
|
||
<taxonomicName id="2BC2A7CA4B1E6ABBE6847C81728ADB33" class="Insecta" family="Scarabaeidae" genus="Prophaenognatha" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Prophaenognatha robusta" order="Coleoptera" pageId="2" pageNumber="55" phylum="Arthropoda" rank="species" species="robusta">Prophaenognatha robusta</taxonomicName>
|
||
<bibRefCitation id="B9D2669222A1344D3715739311EAE4C6" author="Bai, M" journalOrPublisher="Acta Geologica Sinica" pageId="7" pageNumber="60" pagination="984 - 993" title="Prophaenognatha, a new Aclopinae genus from the Yixian Formation, China and its phylogenetic position based on morphological characters (Coleoptera: Scarabaeidae)." url="10.1111/j.1755-6724.2011.00532.x" volume="5" year="2011">Bai et al., 2011</bibRefCitation>
|
||
described in
|
||
<bibRefCitation id="8D6952A2391F0B82D3E17C73EA4F47E6" author="Bai, M" journalOrPublisher="Acta Geologica Sinica" pageId="7" pageNumber="60" pagination="984 - 993" title="Prophaenognatha, a new Aclopinae genus from the Yixian Formation, China and its phylogenetic position based on morphological characters (Coleoptera: Scarabaeidae)." url="10.1111/j.1755-6724.2011.00532.x" volume="5" year="2011">Bai et al. (2011)</bibRefCitation>
|
||
, the Orapa specimen and three outgroup species comprising three families of the
|
||
<taxonomicName id="1F6078FB324A426345450843C7E2B624" lsidName="Hydrophiloidea" pageId="2" pageNumber="55" rank="superfamily" superfamily="Hydrophiloidea">Hydrophiloidea</taxonomicName>
|
||
(
|
||
<taxonomicName id="3DF005EDDB5436B57CD6CE6165A049AC" family="Hydrophilidae" lsidName="" pageId="2" pageNumber="55" rank="family">Hydrophilidae</taxonomicName>
|
||
,
|
||
<taxonomicName id="44CFDBCA579E6AB7B440B518638CDE67" family="Histeridae" lsidName="" pageId="2" pageNumber="55" rank="family">Histeridae</taxonomicName>
|
||
,
|
||
<taxonomicName id="4AAE33CCBFC36929B01F4B903CF1397E" family="Synteliidae" lsidName="" pageId="2" pageNumber="55" rank="family">Synteliidae</taxonomicName>
|
||
). Of the 68 characters in the character set, 27 could be identified with confidence in the Orapa specimen (Table 1). As in
|
||
<bibRefCitation id="0880D9735994E22C9F4CAB9953B7517C" author="Bai, M" journalOrPublisher="Acta Geologica Sinica" pageId="7" pageNumber="60" pagination="984 - 993" title="Prophaenognatha, a new Aclopinae genus from the Yixian Formation, China and its phylogenetic position based on morphological characters (Coleoptera: Scarabaeidae)." url="10.1111/j.1755-6724.2011.00532.x" volume="5" year="2011">Bai et al. (2011)</bibRefCitation>
|
||
, heuristic searches and a bootstrap analysis were performed in Winclada/Nona with the same parameter settings. A strict consensus tree (Fig. 1) was derived from six most parsimonious trees. The phylogeny obtained from the analysis is similar to that in the original study with divisions of higher taxa poorly supported but good support for family groups (see bootstrap values Fig. 1). In the strict consensus tree, the relationship between several families could not be resolved. The Orapa specimen was placed within this polytomy with moderate support as a sister taxa to the
|
||
<taxonomicName id="F49E4DB1003C60610748E48A7DC8FD8F" lsidName="" pageId="2" pageNumber="55" rank="tribe" tribe="Phaenognathini">Phaenognathini</taxonomicName>
|
||
which includes the Yixian fossil
|
||
<taxonomicName id="A1FD3296D8259110CB512D7F9FDB273C" class="Insecta" family="Scarabaeidae" genus="Prophaenognatha" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Prophaenognatha robusta" order="Coleoptera" pageId="2" pageNumber="55" phylum="Arthropoda" rank="species" species="robusta">Prophaenognatha robusta</taxonomicName>
|
||
(Fig. 1). The position of the Orapa fossil in the phylogeny is supported by two character states; antennal club loose, and outer apical angle of protibia simple or slightly produced.
|
||
</paragraph>
|
||
<caption id="6CC908FFAD5DAEFCA90B12F718BD8680" pageId="2" pageNumber="55">
|
||
<paragraph id="D115D9188692882A42A025C727B68A47" pageId="2" pageNumber="55">
|
||
Figure 1. Strict consensus of six most parsimonious trees showing characters (above) and state (below). Bootstrap support values are given in parentheses (tree length=275 steps, ensemble consistency index (CI) = 0.39, ensemble retention index (RI) = 0.64).
|
||
<taxonomicName id="14B9011622E613FF167ADDB036B152C3" class="Insecta" family="Scarabaeidae" genus="Ceafornotensis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Ceafornotensis archratiras" order="Coleoptera" pageId="2" pageNumber="55" phylum="Arthropoda" rank="species" species="archratiras">Ceafornotensis archratiras</taxonomicName>
|
||
is referred to as "Orapa fossil". Character set from
|
||
<bibRefCitation id="E39ECCC2528CB04FB8C9A560072EEC07" author="Bai, M" journalOrPublisher="Acta Geologica Sinica" pageId="7" pageNumber="60" pagination="984 - 993" title="Prophaenognatha, a new Aclopinae genus from the Yixian Formation, China and its phylogenetic position based on morphological characters (Coleoptera: Scarabaeidae)." url="10.1111/j.1755-6724.2011.00532.x" volume="5" year="2011">Bai et al. (2011)</bibRefCitation>
|
||
.
|
||
</paragraph>
|
||
</caption>
|
||
<caption id="C134056E83AAEC1DDC0D9E49C593F794" ID-Table-UUID="C134056E83AAEC1DDC0D9E49C593F794" httpUri="http://table.plazi.org/id/C134056E83AAEC1DDC0D9E49C593F794" pageId="2" pageNumber="55">
|
||
<paragraph id="F7C0FE0E53CBDFDECA4DADCB41DDF2AA" pageId="2" pageNumber="55">
|
||
Table 1. List of 27 characters identified in the Orapa fossil for phylogenetic analysis (Fig. 3). The full characters set of 68 characters is given in
|
||
<bibRefCitation id="A1C0EEB6A7DD5E8D1094B7D57B6F9AC7" author="Bai, M" journalOrPublisher="Acta Geologica Sinica" pageId="7" pageNumber="60" pagination="984 - 993" title="Prophaenognatha, a new Aclopinae genus from the Yixian Formation, China and its phylogenetic position based on morphological characters (Coleoptera: Scarabaeidae)." url="10.1111/j.1755-6724.2011.00532.x" volume="5" year="2011">Bai et al. (2011)</bibRefCitation>
|
||
. Character numbers in the left column refer to the full character set. Characters state for the specimen is given in parentheses in the left column.
|
||
</paragraph>
|
||
</caption>
|
||
<paragraph id="6DE5A345F12D976326D2D2B1690AD78A" pageId="2" pageNumber="55">
|
||
<table id="048BB65C72E3C060F00F3B2060C6734E" pageId="2" pageNumber="55">
|
||
<tr id="CEAB438D56020C74AD61A2408FFA8C6A" pageId="2" pageNumber="55">
|
||
<th id="FC71FE49D21ED3B553E4AC417F0646F5" colspan="1" pageId="2" pageNumber="55" rowspan="1">Character # and state</th>
|
||
<th id="406E7C3C035C82A4D3734811A605C38D" colspan="1" pageId="2" pageNumber="55" rowspan="1">Character and states</th>
|
||
</tr>
|
||
</table>
|
||
</paragraph>
|
||
<paragraph id="35601EFBE12D4B3B0AC7078F802C1DE4" lastPageId="4" lastPageNumber="57" pageId="2" pageNumber="55">
|
||
The close relationship between
|
||
<taxonomicName id="6DC2C4338386DE971688A168F9799DC0" family="Ochodaeidae" lsidName="" pageId="2" pageNumber="55" rank="family">Ochodaeidae</taxonomicName>
|
||
and the
|
||
<taxonomicName id="750C0074C002E7C7CC5D83B63A3D0EEA" family="Hybosoridae" lsidName="" pageId="2" pageNumber="55" rank="family">Hybosoridae</taxonomicName>
|
||
, which is supported in both morphological (
|
||
<bibRefCitation id="2B0397ACD2BACE8C793E08B67647AE1A" author="Browne, J" journalOrPublisher="Systematic Entomology" pageId="7" pageNumber="60" pagination="51 - 84" title="A phylogeny of the families of Scarabaeoidea (Coleoptera)." url="10.1046/j.1365-3113.1999.00067.x" volume="24" year="1999">Browne and Scholtz 1999</bibRefCitation>
|
||
;
|
||
<bibRefCitation id="551493EA99D2D70321ED96717C9D3D0B" author="Scholtz, CH" journalOrPublisher="Journal of the Entomological Society of Southern Africa" pageId="9" pageNumber="62" pagination="207 - 240" title="Phylogeny and systematics of the Ochodaeidae (Insecta: Coleoptera: Scarabaeoidea)." volume="51" year="1988">Scholtz et al. 1988</bibRefCitation>
|
||
) and
|
||
<pageBreakToken id="BB8DEF969A7BD627F2238F170354D6D2" pageId="3" pageNumber="56" start="start">molecular-based</pageBreakToken>
|
||
phylogenies (
|
||
<bibRefCitation id="A441B577F11FA8FB1B44D342EC652F08" author="Ahrens, D" journalOrPublisher="Proceedings of the Royal Society of London B: Biological Sciences" pageId="7" pageNumber="60" title="The evolution of scarab beetles tracks the sequential rise of angiosperms and mammals." url="10.1098/rspb.2014.1470" volume="281" year="2014">Ahrens et al. 2014</bibRefCitation>
|
||
;
|
||
<bibRefCitation id="BF95E769146DCBC893C85825CDA4374F" author="Smith, ABT" journalOrPublisher="Coleopterists Society Monograph" pageId="9" pageNumber="62" pagination="35 - 46" title="An overview of the classification and evolution of the major scarab beetle clades (Coleoptera: Scarabaeoidea) based on preliminary molecular analyses." url="10.1649/0010-065X(2006)60[35:AOOTCA]2.0.CO;2" volume="5" year="2006">Smith et al. 2006</bibRefCitation>
|
||
), was not apparent in the strict consensus tree and may suggest that the placement of
|
||
<taxonomicName id="269785042DE8600ED51BBC13FBBA76B5" family="Ochodaeidae" lsidName="" pageId="3" pageNumber="56" rank="family">Ochodaeidae</taxonomicName>
|
||
as a sister taxa to the
|
||
<taxonomicName id="2072EDF860935EFDCC16FC2A182B2EA7" lsidName="" pageId="3" pageNumber="56" rank="tribe" tribe="Phaenognathini">Phaenognathini</taxonomicName>
|
||
is erroneous. The
|
||
<taxonomicName id="41B7C21B7D7344F3F241E276CC7DECC5" lsidName="" pageId="3" pageNumber="56" rank="tribe" tribe="Phaenognathini">Phaenognathini</taxonomicName>
|
||
, recently established by
|
||
<bibRefCitation id="052F598417C3620FDFA854B120E17E18" author="Ocampo, FC" journalOrPublisher="Zootaxa" pageId="8" pageNumber="61" pagination="1 - 29" title="Revision of the scarab family Aclopinae Blanchard (Coleoptera: Scarabaeidae) in Argentina and Chile." volume="3409" year="2012">Ocampo and Mondaca (2012)</bibRefCitation>
|
||
, contains the genera
|
||
<taxonomicName id="5892DD00167E20C5724C2DB2C10BB7C9" class="Insecta" family="Scarabaeidae" genus="Phaenognatha" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Phaenognatha" order="Coleoptera" pageId="3" pageNumber="56" phylum="Arthropoda" rank="genus">Phaenognatha</taxonomicName>
|
||
Hope, 1845 and
|
||
<taxonomicName id="0B57A35174A7114945D57589661C42C5" class="Insecta" family="Scarabaeidae" genus="Neophaenognatha" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Neophaenognatha" order="Coleoptera" pageId="3" pageNumber="56" phylum="Arthropoda" rank="genus">Neophaenognatha</taxonomicName>
|
||
Allsopp, 1983 formerly placed within the
|
||
<taxonomicName id="165889127A2B000AF566E1E81951AAA0" lsidName="" pageId="3" pageNumber="56" rank="subfamily" subfamily="Aclopinae">Aclopinae</taxonomicName>
|
||
. The position of the
|
||
<taxonomicName id="D584276D660BC8EF7EFAA8FFA4D434D9" lsidName="" pageId="3" pageNumber="56" rank="tribe" tribe="Phaenognathini">Phaenognathini</taxonomicName>
|
||
within the
|
||
<taxonomicName id="34A4F85615765E22151BE4EC131E3FF8" lsidName="" pageId="3" pageNumber="56" rank="subfamily" subfamily="Melolonthinae">Melolonthinae</taxonomicName>
|
||
remains unclear although
|
||
<bibRefCitation id="623135F4CA41F418930836630AB826DD" author="Ocampo, FC" journalOrPublisher="Zootaxa" pageId="8" pageNumber="61" pagination="1 - 29" title="Revision of the scarab family Aclopinae Blanchard (Coleoptera: Scarabaeidae) in Argentina and Chile." volume="3409" year="2012">Ocampo and Mondaca (2012)</bibRefCitation>
|
||
have hypothesised a relationship between the
|
||
<taxonomicName id="213ACA417E3B6427FC0D2BDB42444596" lsidName="" pageId="3" pageNumber="56" rank="tribe" tribe="Phaenognathini">Phaenognathini</taxonomicName>
|
||
and the
|
||
<taxonomicName id="31AAB0822BC8E657B9AD5D0EB9F16250" lsidName="" pageId="3" pageNumber="56" rank="tribe" tribe="Lichniini">Lichniini</taxonomicName>
|
||
suggesting that they may "constitute a distinct and ancient lineage of
|
||
<taxonomicName id="466D955B603D65CCB1165B0E83C7839F" family="Scarabaeidae" lsidName="" pageId="3" pageNumber="56" rank="family">Scarabaeidae</taxonomicName>
|
||
". Evidence from a preliminary molecular analysis (
|
||
<bibRefCitation id="6809F6C15BE34884A466884C5402EA29" author="Smith, ABT" journalOrPublisher="Coleopterists Society Monograph" pageId="9" pageNumber="62" pagination="35 - 46" title="An overview of the classification and evolution of the major scarab beetle clades (Coleoptera: Scarabaeoidea) based on preliminary molecular analyses." url="10.1649/0010-065X(2006)60[35:AOOTCA]2.0.CO;2" volume="5" year="2006">Smith et al. 2006</bibRefCitation>
|
||
)
|
||
<pageBreakToken id="6BD92126C9226355130A619ED3216DBB" pageId="4" pageNumber="57" start="start">indicates</pageBreakToken>
|
||
that several smaller taxa belong within a
|
||
<taxonomicName id="866BB319DEAE7DD6A4C158AB118D0E95" lsidName="" pageId="4" pageNumber="57" rank="subfamily" subfamily="Melolonthinae">Melolonthinae</taxonomicName>
|
||
clade or
|
||
<normalizedToken id="F1E50AE19FE24507727D3AE20DCF8F10" originalValue="‘grade’">'grade'</normalizedToken>
|
||
. These taxa include
|
||
<taxonomicName id="1F40C8269A3701FE69E543AFCD404C5C" lsidName="" pageId="4" pageNumber="57" rank="tribe" tribe="Lichniini">Lichniini</taxonomicName>
|
||
, removed from the
|
||
<taxonomicName id="194DCEED28B8EEA17ECB13A3DE18B98A" family="Glaphyridae" lsidName="" pageId="4" pageNumber="57" rank="family">Glaphyridae</taxonomicName>
|
||
(
|
||
<bibRefCitation id="54CCB4A60901985EC33E11C4A271AB8E" author="Hawkins, SJ" journalOrPublisher="Zootaxa" pageId="7" pageNumber="60" pagination="1 - 63" title="A revision of the Chilean tribe Lichniini Burmeister, 1844 (Coleoptera: Scarabaeidae: Melolonthinae)." volume="1266" year="2006">Hawkins 2006</bibRefCitation>
|
||
), and subfamilies of the
|
||
<taxonomicName id="2C48879858E72F3A458F3F091945F76D" family="Scarabaeidae" lsidName="" pageId="4" pageNumber="57" rank="family">Scarabaeidae</taxonomicName>
|
||
;
|
||
<taxonomicName id="C9219BBCD83482053D662343DDFBAC84" lsidName="" pageId="4" pageNumber="57" rank="subfamily" subfamily="Aclopinae">Aclopinae</taxonomicName>
|
||
,
|
||
<taxonomicName id="5362CB99D6A60814D85C17AA9EC5434E" lsidName="" pageId="4" pageNumber="57" rank="subfamily" subfamily="Allidiostomatinae">Allidiostomatinae</taxonomicName>
|
||
,
|
||
<taxonomicName id="294D52F0AAA7AFB515E65B44B2A6CD77" lsidName="" pageId="4" pageNumber="57" rank="subfamily" subfamily="Euchirinae">Euchirinae</taxonomicName>
|
||
,
|
||
<taxonomicName id="F40EBCAE62FD2FF9756329A91F633677" lsidName="" pageId="4" pageNumber="57" rank="subfamily" subfamily="Orphninae">Orphninae</taxonomicName>
|
||
and
|
||
<taxonomicName id="4B929B578E469DBF7B3156A81643AFE1" lsidName="" pageId="4" pageNumber="57" rank="subfamily" subfamily="Pachypodinae">Pachypodinae</taxonomicName>
|
||
. The paraphyletic grade of the
|
||
<taxonomicName id="4F65BE7047724DBECACA06F655C30AA9" lsidName="" pageId="4" pageNumber="57" rank="subfamily" subfamily="Melolonthinae">Melolonthinae</taxonomicName>
|
||
is also supported by recent studies (
|
||
<bibRefCitation id="B48764E8073D838E55DC05D3F0537D71" author="Ahrens, D" journalOrPublisher="Proceedings of the Royal Society of London B: Biological Sciences" pageId="7" pageNumber="60" title="The evolution of scarab beetles tracks the sequential rise of angiosperms and mammals." url="10.1098/rspb.2014.1470" volume="281" year="2014">Ahrens et al. 2014</bibRefCitation>
|
||
;
|
||
<bibRefCitation id="E922EB94BBCDB1398CC080158EF91EA7" author="McKenna, DD" journalOrPublisher="Systematic Entomology" pageId="8" pageNumber="61" pagination="35 - 60" title="Phylogeny and evolution of Staphyliniformia and Scarabaeiformia: forest litter as a stepping stone for diversification of nonphytophagous beetles." url="10.1111/syen.12093" volume="40" year="2015">McKenna et al. 2015</bibRefCitation>
|
||
). Given the uncertainties of the current phylogeny, the Orapa specimen is placed as
|
||
<taxonomicName id="C84AEC83BAA96DB5206637B6F81057CF" lsidName="" pageId="4" pageNumber="57" rank="subfamily" subfamily="Melolonthinae">Melolonthinae</taxonomicName>
|
||
incertae sedis.
|
||
</paragraph>
|
||
</subSubSection>
|
||
</treatment>
|
||
</document> |